A novel gene Bractea (bra) controls the formation of an indeterminate bractless inflorescence in Arabidopsis thaliana

The morphological and genetic studies of the bra mutant of Arabidopsis thaliana (L.) Heynh. from the collection of the Department of Genetics and Breeding, Moscow State University, showed that the BRA gene controls the main stages of inflorescence development: it suppresses the development of leaflike organs subtending flowers (bracts) and inhibits the formation of the terminal flower. Inactivation of the BRA gene leads to the transition from the indeterminate bractless inflorescence characteristic of the family Cruciferaceae to the determinate bracteose inflorescence. The BRA gene plays a regulatory role and was probably involved in the conversion of the bracteose determinate inflorescence to the bractless indeterminate inflorescence during the origin of ancestral crucifers.     

A NewBRACTEA(BRA) Gene Controlling the Formation of an Indeterminate Bractless Inflorescence in Arabidopsis thaliana

The morphological and genetic studies of the bramutant of Arabidopsis thaliana(L.) Heynh. from the collection of the Department of Genetics and Breeding, Moscow State University, showed that the BRAgene controls the major stages of inflorescence development: it suppresses the development of leaves subtending flowers (bracts) and inhibits the formation of the terminal flower. Inactivation of the BRAgene leads to the transition from the indeterminate bractless inflorescence characteristic of the family Cruciferaceae to the determinate bracteose inflorescence. It is suggested that the BRAgene is a regulator gene probably involved in the conversion of the bracteose determinate inflorescence to the indeterminate ebracteate inflorescence during the origin of ancestral crucifers.     

ABERRANT PANICLE ORGANIZATION 2/RFL, the rice ortholog of Arabidopsis LEAFY, suppresses the transition from inflorescence meristem to floral meristem through interaction with APO1

The temporal and spatial control of meristem identity is a key element in plant development. To better understand the molecular mechanisms that regulate inflorescence and flower architecture, we characterized the rice aberrant panicle organization 2 (apo2) mutant which exhibits small panicles with reduced number of primary branches due to the precocious formation of spikelet meristems. The apo2 mutants also display a shortened plastochron in the vegetative phase, late flowering, aberrant floral organ identities and loss of floral meristem determinacy. Map-based cloning revealed that APO2 is identical to previously reported RFL gene, the rice ortholog of the Arabidopsis LEAFY (LFY) gene. Further analysis indicated that APO2/RFL and APO1, the rice ortholog of Arabidopsis UNUSUAL FLORAL ORGANS, act cooperatively to control inflorescence and flower development. The present study revealed functional differences between APO2/RFL and LFY. In particular, APO2/RFL and LFY act oppositely on inflorescence development. Therefore, the genetic mechanisms for controlling inflorescence architecture have evolutionarily diverged between rice (monocots) and Arabidopsis (eudicots).     

珙桐(Davidia involucrata Baill.)苞片功能性状及性状间关系对海拔的响应

珙桐(Davidia involucrata Baill.)的苞片被认为是对传粉者和非生物因素等驱动力的适应,往往受到环境因子的影响。为揭示其功能性状及其性状间关系对海拔的响应,本研究采用独立样本t检验和标准化主轴分析方法对比了四川龙苍沟1400和1800 m不同海拔珙桐种群中花苞片的功能性状和性状间关系。结果显示:1)低海拔(1400 m)珙桐种群大、小苞片的长、宽和面积,以及单花苞片总面积均显著大于高海拔(1800 m)种群(P<0.05),且高、低海拔种群的大、小苞片面积及重量呈等比例生长; 2)低海拔珙桐种群大、小苞片干重及单花苞片总干重均显著高于高海拔种群(P<0.05); 3)高、低海拔种群的花序轴长与干重均无显著差异(P=0.446; P=0.791),高海拔花序轴干重与长间呈异速生长关系,而低海拔呈等速生长关系; 4)珙桐大、小苞片的长宽间、面积间以及重量间关系在高、低海拔上均表现出等速生长,单花苞片的总面积和总干重分别与花序轴干重在高、低海拔上均表现出等速生长关系,而与花序轴长表现出异速生长关系。上述结果表明,珙桐苞片的功能性状及其性状间关系在不同海拔存在...     

The Morphology of the Inflorescence in Ranunculus bulbosus L.

This paper records the form variation of 750 inflorescencesof Ranunculus bulbosus L. collected randomly from each of twolarge colonies growing on permanent grassland. Each inflorescence has a terminal flower, 1–4 bracts onthe main axis and up to 8 flowers borne on cymes subtended bythese bracts. Over 75 per cent, of each sample consists of inflorecenceswith 2 or 3 bracts on the main axis and 2–4 flowers. Thenumber of flowers increases with the number of bracts on themain axis and evidence is given that the 4-bract 9-floweredinflorescence may be nearly the largest and most complex thatcan be produced under these conditions. The distribution of flowers in the axillary cymes is such thatthe inflorescences tend to be radially symmetrical and pyramidalin form. This is so even thought with increase in the numberof bracts on the main axis the proportion of axillary flowersdecreases in the lowest cyme and increases in the cyme above. It is considered that the form and size of the inflorescencecan be related to the vigour of the plant and to the mechanicaland nutritional problems involved. A comparison of the varioustypes of inflorescences found probably reflects the developmentalsequence of flower production. It also indicates that thereis competition between certain potential flower positions asthe inflorescence develops.     

Arabidopsis thaliana (L.) Heynh. as a model object for studying genetic control of morphogenesis

A vast amount of information on the genetic control of plant development has been obtained in Arabidopsis thaliana with classical genetic and molecular biological methods. The genes involved in multistep regulation of floral morphogenesis have been identified. The formation of floral meristem is controlled by the LEAFY (LFY), UNUSUAL FLORAL ORGANS (UFO), APETALA1 (AP1), and APETALA2 (AP2) genes. Studies of the abruptus and bractea recessive monogenic mutants from the collection of the Department of Genetics and Selection, Moscow State University, showed that the ABRUPTUS (ABR) and BRACTEA (BRA) genes also play an important role in inflorescence differentiation. The ABR gene controls the early formation of organ primordia on the inflorescence and the formation of floral organ primordia after floral initiation. Further differentiation of inflorescence organ primordia in vegetative or generative organs depends on the activity of the LFY gene, and floral organ identity is determined by the homeotic genes. Presumably, the major function of the ABR gene is to determine the auxin polar transport. The BRA gene suppresses the development of bracts on the inflorescence and constrains cell division at the base of primordia of rosette and cauline leaves.     

DEVELOPMENTAL STUDIES IN PTYCHOSPERMA (PALMAE). I. THE INFLORESCENCE AND THE FLOWER CLUSTER

This paper describes inflorescence structure, including organogenesis of the panicle and flower clusters and vasculature of flowering branches, for two species of Ptychosperma, a genus of arecoid palms. The inflorescence is an infrafoliar panicle with up to four orders of branches in a spirodistichous arrangement conforming to an irregular one-half phyllotaxy. The primordium of the inflorescence is crescentic and the apex has two tunica layers, a group of central cells, and a rib meristem. The distal flower-bearing parts or rachillae of all branches develop acropetally early in ontogeny and are vertically oriented in the bud. Although these rachillae terminate branches of different sizes and orders, they are similar in size and in number of flower clusters produced. Internodes and lower parts of branches develop later. Bracts of four types are produced: a prophyll and empty peduncular bract, bracts which subtend lateral branches, bracts subtending triads, and floral bracteoles. The prophyll and peduncular bracts are tubular and completely closed around all branches until about three months before the flowers reach anthesis. Bracts subtending lateral branches and those that subtend triads enlarge by small amounts of apical, adaxial, and marginal growth to cover subtended apices during early ontogeny, but are small to absent at maturity. Flower clusters are triads of two lateral staminate and a central pistillate flower. Organogenesis indicates that the triad is a sympodial unit. Flowers develop successively, each floral apex bearing a bracteole that subtends the next flower. The vasculature of the inflorescence may be divided into two systems. Bundles of the main axis extend acropetally into the vertically oriented branches as they are initiated and form a central cylinder of larger bundles in each branch. Flower clusters are supplied by a peripheral system of smaller bundles that develop later in relation to the developing floral organs. Bundles of the peripheral system branch frequently, but branching levels are irregular. The irregular branching of peripheral bundles appears related to the phyllotaxy of the flower clusters and the random right or left position of the first flower of the triad. The level of branching of a bundle may depend on the position of a floral primordium with respect to an existing procambial strand. Three (-4) bundles supply each staminate flower and six (-10) the pistillate flower. The histologically specialized inflorescence has stomata and contains abundant starch. Tannins and raphides, spherical silica bodies, and various forms of sclerenchyma appear in sequence and apparently provide support and protection during the long exposure of the branches.     

DFL, a FLORICAULA/LEAFY homologue gene from Dendranthema lavandulifolium is expressed both in the vegetative and reproductive tissues

FLO/LFY homologue genes were initially characterized as floral meristem identity genes and play a key role in flower development among diverse species. The inflorescence organization of chrysanthemum differs from typical dicotyledons such as Arabidopsis and Antirrhinum as clear sepals are absent, and instead, a pappus, a rudimentary sepal, is formed. To understand the mechanism of reproduction of chrysanthemum at the molecular level, DFL, a FLORICAULA/LEAFY homologous gene, was cloned from Dendranthema lavandulifolium, which is one of the original species of chrysanthemum. The DFL gene consists of a 1,236-bp open reading frame and encodes a putative protein of 412 amino acids, which is 63% identical to LFY and 70% to FLO. The expression patterns of DFL during the flower development were analyzed, and RT-PCR results showed that DFL was strongly expressed in the flower bud. In situ hybridization experiments showed that it is strongly expressed in the inflorescence bract, petal and stamen primordial tissues throughout the inflorescence development. Its expression signals were also detected in stems, leaf primordial tissues and developing inflorescence bracts.     

MISSING FLOWERS gene controls axillary meristems initiation in sunflower

The initiation and growth of axillary meristems are fundamental components of plant architecture. Here, we describe the mutant missing flowers (mf) of Helianthus annuus characterized by the lack of axillary shoots. Decapitation experiments and histological analysis indicate that this phenotype is the result of a defect in axillary meristem initiation. In addition to shoot branching, mutation affects floral differentiation. The indeterminate inflorescence of sunflower (capitulum) is formed of a large flat meristem which produces floret primordia in multiple spirals. In wildtype plants a bisecting crease divides each primordium in two distinct bumps that adopt different fate. The peripheral (abaxial) part of the primordium becomes a small leaf-like bract and the adaxial part becomes a flower. In the mf mutant, the formation of flowers at the axil of bracts is precluded. Histological analyses show that in floret primordia of the mutant a clear subdivision in dyads is not established. The primordia progressively bend inside and only large involucral floral bracts are developed. The results suggest that the MISSING FLOWERS gene is essential to provide or perceive an appropriate signal to the initiation of axillary meristems during both vegetative and reproductive phases.     

LEAFY and the evolution of rosette flowering in violet cress (Jonopsidium acaule, Brassicaceae)

Arabidopsis and most other Brassicaceae produce an elongated inflorescence of mainly ebracteate flowers. However, the early-flowering species violet cress (Jonopsidium acaule) and a handful of other species produce flowers singly in the axils of rosette leaves. In Arabidopsis the gene LEAFY (LFY) is implicated in both the determination of flower meristem identity and in the suppression of leaves (bracts) that would otherwise subtend the flowers. In this study we examined the role of LFY homologs in the evolution of rosette flowering in violet cress. We cloned two LFY homologs, vcLFY1 and vcLFY2, from violet cress. Their exon sequences show ～90% nucleotide similarity with Arabidopsis LFY and 99% similarity to each other. We used in situ hybridization to study vcLFY expression in violet cress. The patterns were very similar to LFY in Arabidopsis except for stronger expression in the shoot apical meristem outside of the region of flower meristem initiation. It is possible that the relatively diffuse expression of vcLFY contributes to the lack of bract suppression in violet cress. Additionally, the earliest flowers produced by violet cress express vcLFY, suggesting that accelerated flowering in violet cress could also result from changes in the regulation of vcLFY.     

Regulatory networks that function to specify flower meristems require the function of homeobox genes PENNYWISE and POUND-FOOLISH in Arabidopsis

Flowering is a major developmental phase change that transforms the fate of the shoot apical meristem (SAM) from a leaf-bearing vegetative meristem to that of a flower-producing inflorescence meristem. In Arabidopsis, floral meristems are specified on the periphery of the inflorescence meristem by the combined activities of the FLOWERING LOCUS T (FT)–FD complex and the flower meristem identity gene, LEAFY ( LFY ). Two redundant functioning homeobox genes, PENNYWISE ( PNY ) and POUND-FOOLISH ( PNF ), which are expressed in the vegetative and inflorescence SAM, regulate patterning events during reproductive development, including floral specification. To determine the role of PNY and PNF in the floral specification network, we characterized the genetic relationship of these homeobox genes with LFY and FT . Results from this study demonstrate that LFY functions downstream of PNY and PNF. Ectopic expression of LFY promotes flower formation in pny pnf plants, while the flower specification activity of ectopic FT is severely attenuated. Genetic analysis shows that when mutations in pny and pnf genes are combined with lfy , a synergistic phenotype is displayed that significantly reduces floral specification and alters inflorescence patterning events. In conclusion, results from this study support a model in which PNY and PNF promote LFY expression during reproductive development. At the same time, the flower formation activity of FT is dependent upon the function of PNY and PNF.     

杏黄兜兰的花发育过程及引种栽培

通过形态解剖研究了杏黄兜兰花的发育过程,发现杏黄兜兰5月份发育出花序原基,6～7月份发育出花序鞘,7～8月份发育出苞片,8月底至9月完成花器官形态分化过程。形态分化完成后杏黄兜兰的花器官继续快速增长。退化雄蕊、能育雄蕊、花柱在花形成后早期不融合,在后期才融合形成合蕊柱,且因为相对生长速率的不同,三者的相对位置、形态也发生改变。此外,在7～9月份引种至昆明栽培的不同批次杏黄兜兰中,产生花芽比率显著不同,进一步证实8月底至9月初是杏黄兜兰进行花器官形态分化的关键时期。     

ONTOGENY OF THE INFLORESCENCE OF SAURURUS CERNUUS (SAURURACEAE)

The spicate inflorescence of Saururus cernuus L. (Saururaceae) results from the activity of an inflorescence apical meristem which produces 200–300 primordia in acropetal succession. The inflorescence apex arises by conversion of the terminal vegetative apex. During transition the apical meristem increases greatly in height and width and changes its cellular configuration from one of tunica-corpus to one of mantle (with two tunica layers) and core. Primordia are initiated by periclinal divisions in the subsurface layer. These are “common” primordia, each of which subsequently divides to produce a floral apex above and a bract primordium below. The bract later elongates so that the flower appears borne on the bract. All common primordia are formed by the time the inflorescence is about 4.4 mm long; the apical meristem ceases activity at this stage. As cessation approaches, cell divisions become rare in the apical meristem, and height and width of the meristem above the primordia diminish, as primordia continue to be initiated on the flanks. Cell differentiation proceeds acropetally into the apical meristem and reaches the summital tunica layers last of all. Solitary bracts are initiated just before apical cessation, but no imperfect or ebracteate flowers are produced in Saururus. The final event of meristem activity is hair formation by individual cells of the tunica at the summit, a feature not previously reported for apical meristems.     

Development and morphology of flowers and inflorescences in Balanophora papuana and B. elongata (Balanophoraceae)

Extreme modification and reduction in floral morphology presents an obstacle to determining the evolutionary relationships and homologies of the holoparasites in Balanophoraceae. Developing flowers and inflorescences of two dioecious species, Balanophora papuana and B. elongata, were compared to each other and to the monoecious B. fungosa. Intermingled with flowers in the male inflorescences are bracts (B. elongata) or bract parts (B. papuana). In the latter, early cessation of bract tip growth results in two half-bracts, which become displaced during inflorescence elongation, thus disproving the view that these bract-like structures are axial in nature. Male flower primordia emerge in positions axillary to the dividing bracts, and both arise in a spiral sequence. This pattern is modified in B. papuana by the formation of pseudowhorls of four. In both species, the staminate flowers consist of a generally four-merous perianth and a synandrium of congenitally fused stamens. Male flower and bract ontogeny (but not pollen sacs) conform to patterns seen in other angiosperms. More problematic are the carpellate flowers whose primordia arise in irregular order between club-shaped, radially symmetrical organs called claviform bodies. The interpretation that these bodies are homologous to the peltate bracts of Helosideae appears plausible, but cannot explain their nonspiral initiation and radial symmetry.     

The role of teosinte glume architecture (tga1) in coordinated regulation and evolution of grass glumes and inflorescence axes

? Hardened floral bracts and modifications to the inflorescence axis of grasses have been hypothesized to protect seeds from predation and/or aid seed dispersal, and have evolved multiple times independently within the family. Previous studies have demonstrated that mutations in the maize (Zea mays ssp. mays) gene teosinte glume architecture (tga1) underlie a reduction in hardened structures, yielding free fruits that are easy to harvest. It remains unclear whether the causative mutation(s) occurred in the cis-regulatory or protein-coding regions of tga1, and whether similar mutations in TGA1-like genes can explain variation in the dispersal unit in related grasses. ? To address these questions TGA1-like genes were cloned and sequenced from a number of grasses and analyzed phylogenetically in relation to morphology; protein expression was investigated by immunolocalization. ? TGA1-like proteins were expressed throughout the spikelet in the early development of all grasses, and throughout the flower of the grass relative Joinvillea. Later in development, expression patterns differed between Tripsacum dactyloides, maize and teosinte (Z. mays ssp. parviglumis). ? These results suggest an ancestral role for TGA1-like genes in early spikelet development, but do not support the hypothesis that TGA1-like genes have been repeatedly modified to affect glume and inflorescence axis diversification.     

A theory for inflorescence development and flower formation based on morphological and biophysical analysis in Echeveria

Floral development is generally viewed as involving interactions between recently made organs and generative activity on the apical dome; one set of floral organs is thought to induce the next. To investigate such interactions, flowering in Echeveria derenbergii (J. Purpus) was studied at two levels of structure. At the larger, morphological, level the inflorescence apex is shown to have simple cyclic development. Seen from above, it elongates horizontally, then forms a transverse cleft to demarcate a flower primordium in one of two rows. The meristem then elongates at 90° to its previous axis, also horizontally, and demarcates a flower in the other row. Activity on the apical surface correlates well with the nature and activity of adjacent sub-apical organs. For example, the 90° shifts in elongation of the meristem correlate with that tissue's being attached, laterally, to successive large growing bracts whose bases lie at 90°. Also, on the flower primordium, the five sepals arise in a spiral sequence which correlates with one of increasing age, since formation by the cleft, of the edges of the primordium.The second level of study was to test whether the developmental correlations could have a biophysical explanation. By biophysical theory, organs arise where the dome surface is structurally predisposed to bulge. This is a function of the cellulose reinforcement pattern in the surface. Successive patterns of cellulose reinforcement in isolated surface layers from floral organs were determined using polarized light. This was done for the cyclic activity of the inflorescence meristem and the development of the flower. The results indicate that patterns of cellulose reinforcement on the apical dome surface could lead to the production of organs, through local promotion of bulging of the tunica. Subsequent growth of the base of each organ stretches the adjacent dome tissue in a directional fashion. Cytoskeletal responses of these stretched cells lead to new cellulose alignments on the dome which generate the reinforcement pattern for the next round of organs.Abbreviations F floral meristem tissue which will directly produce a flower, starting with sepals - I inflorescence meristem tissue, generally oval in top view and bounded by two bracts, that produces both floral tissue (F) and additional I meristem tissue - I-max the maximal size of I tissue before it bifurcates into F tissue and I tissue (I-min) - I-min the minimal size of I tissue just after it has bifurcated to produce F tissue and I tissue     

MADS-box gene expression and implications for developmental origins of the grass spikelet

Basic questions regarding the origin and evolution of grass (Poaceae) inflorescence morphology remain unresolved, including the developmental genetic basis for evolution of the highly derived outer spikelet organs. To evaluate homologies between the outer sterile organs of grass spikelets and inflorescence structures of nongrass monocot flowers, we describe expression patterns of APETALA1/FRUITFULL-like (AP1/FUL) and LEAFY HULL STERILE-like (LHS1) MADS-box genes in an early-diverging grass (Streptochaeta angustifolia) and a nongrass outgroup (Joinvillea ascendens). AP1/FUL-like genes are expressed only in floral organs of J. ascendens, supporting the hypothesis that they mark the floral boundary in nongrass monocots, and JaLHS1/OsMADS5 is expressed in the inner and outer tepals, stamen filaments and pistil. In S. angustifolia, SaFUL2 is expressed in all 11 (or 12) bracts of the primary inflorescence branch, but not in the suppressed floral bract below the abscission zone. In contrast, SaLHS1 is only expressed in bracts 6-11 (or 12). Together, these data are consistent with the hypotheses that (1) bracts 1-5 of S. angustifolia primary inflorescence branches and glumes of grass spikelets are homologous and that (2) the outer tepals of immediate grass relatives, bracts 6-8 of S. angustifolia, and the lemma/palea are homologous, although other explanations are possible.